Key Points
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Endospores are formed by Bacillus, Clostridium and close relatives, generally in response to nutritional stress. The process of endospore formation is important both ecologically and practically, and it represents a simple, experimentally tractable example of cellular development and differentiation.
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Several hundred genes are specifically devoted to sporulation and the general regulatory pathways — involving five sigma factors and numerous other transcriptional regulators — are now well understood.
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Sporulation begins with a modified asymmetric cell division. The cell-division machinery is moved from its usual mid-cell position, to sites near each of the cell poles. One subpolar site is chosen for division. This generates the small prespore and the larger mother-cell compartments. Chromosome segregation into the prespore occurs, unusually, after septation, requiring a DNA transport protein, SpoIIIE.
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Compartment-specific transcription is initiated by cell-specific activation of two different sigma factors: σF in the prespore and σE in the mother cell. Complex regulatory mechanisms regulate the activity of these sigma factors so that they are activated at the right time and in the right compartment. σF becomes active first, by regulatory mechanisms that are responsive to formation of the sporulation septum.
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Soon after septation, the cell-wall material in the septum is hydrolysed and the membranes of the septum migrate around the prespore, resulting in complete enclosure of the prespore by the mother-cell cytoplasm. This engulfment process generates the unique topological state by which endospores differ from all other spores.
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Late-gene expression in the two compartments is driven by σG (prespore) and σK (mother cell). These factors turn on new sets of genes that encode the proteins that are needed for formation of the protective cortex and spore-coat structures, and they bring about the massive physiological and biochemical changes that occur in the spore and that produce its unique resistance and dormancy properties.
Abstract
Spore formation in bacteria poses a number of biological problems of fundamental significance. Asymmetric cell division at the onset of sporulation is a powerful model for studying basic cell-cycle problems, including chromosome segregation and septum formation. Sporulation is one of the best understood examples of cellular development and differentiation. Fascinating problems posed by sporulation include the temporal and spatial control of gene expression, intercellular communication and various aspects of cell morphogenesis.
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Acknowledgements
I apologise to colleagues whose work has not been cited in full owing to space constraints. I thank A. Feucht for helpful comments on the manuscript. Work in the Errington laboratory is supported by grants from the Biotechnology and Biological Sciences Research Council, the Medical Research Council and the Human Frontier Science Program.
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Glossary
- PRESPORE
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Equivalent to the forespore but sometimes used specifically for the small compartment before completion of engulfment.
- FORESPORE
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A small compartment that is formed after asymmetric division. It is sometimes used specifically for the small compartment specifically after completion of engulment.
- MOTHER CELL
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The large compartment in which the spore develops.
- SPORANGIUM
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The literal meaning is equivalent to the mother cell, but it is frequently used to refer to the two-compartment sporulating organism (that is, the prespore/forespore plus the mother cell).
- SIGMA FACTOR
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A subunit of the RNA polymerase holoenzyme that is required for promoter sequence recognition and the ability to initiate transcription.
- TUBULIN
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A eukaryotic cytoskeletal protein that is used to form microtubules.
- ANTI-SIGMA FACTOR
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A negative transcriptional regulator that acts by binding to a sigma factor and preventing its activity. An anti-anti-sigma factor, in turn, counteracts the action of an anti-sigma factor.
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Errington, J. Regulation of endospore formation in Bacillus subtilis. Nat Rev Microbiol 1, 117–126 (2003). https://doi.org/10.1038/nrmicro750
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DOI: https://doi.org/10.1038/nrmicro750
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